Forest reclamation of fly ash deposit: a field study on appraisal of mycorrhizal inoculation

An extensive field trial was established on a fly ash deposit (1) to evaluate whether the inoculation with arbuscular mycorrhizal fungi (AMF) and/or ectomycorrhizal fungi (EcMF) improves growth and survival of 13 planted tree species and (2) to trace the inoculated mycorrhizal fungi in tree roots after one growing season. Molecular methods were applied to characterize AMF and EcMF entering the studied system (inocula, native soil, and roots of nursery seedlings). Biometric parameters and mortality of the trees were recorded and the presence of AMF and EcMF in sampled trees was determined both microscopically and genetically. Mycorrhizal inoculation did not improve survival or growth of any tree species. Most AMF‐host and all EcMF‐host seedlings were highly precolonized already from the nursery. An abundant and diverse AMF community was also found in the field soil. The AMF inoculum taxa partially overlapped with AMF in the native soil and in the precolonized roots. After one season, the only two inoculum‐unique AMF taxa were detected in host species non‐precolonized or only partially precolonized from the nursery. The components of EcMF inoculum were not detected in any sampled tree. After the season, the ectomycorrhizal hosts maintained most of their original EcMF taxa gathered in nursery, some tree species were additionally colonized by EcMF probably originating from the soil. Our results show considerable self‐restoration potential of nature on the target site. Mycorrhizal inoculation thus did not bring any conclusive advantage to the planted trees and seems superfluous for reclamation practice on the fly ash deposit.     

Comparison of root-associated communities of native and non-native ectomycorrhizal hosts in an urban landscape

Non-native tree species are often used as ornamentals in urban landscapes. However, their root-associated fungal communities remain yet to be examined in detail. Here, we compared richness, diversity and community composition of ectomycorrhizosphere fungi in general and ectomycorrhizal (EcM) fungi in particular between a non-native Pinus nigra and a native Quercus macrocarpa across a growing season in urban parks using 454-pyrosequencing. Our data show that, while the ectomycorrhizosphere community richness and diversity did not differ between the two host, the EcM communities associated with the native host were often more species rich and included more exclusive members than those of the non-native hosts. In contrast, the ectomycorrhizosphere communities of the two hosts were compositionally clearly distinct in nonmetric multidimensional ordination analyses, whereas the EcM communities were only marginally so. Taken together, our data suggest EcM communities with broad host compatibilities and with a limited numbers of taxa with preference to the non-native host. Furthermore, many common fungi in the non-native Pinus were not EcM taxa, suggesting that the fungal communities of the non-native host may be enriched in non-mycorrhizal fungi at the cost of the EcM taxa. Finally, while our colonization estimates did not suggest a shortage in EcM inoculum for either host in urban parks, the differences in the fungi associated with the two hosts emphasize the importance of using native hosts in urban environments as a tool to conserve endemic fungal diversity and richness in man-made systems.     

High richness of ectomycorrhizal fungi and low host specificity in a coastal sand dune ecosystem revealed by network analysis

Ectomycorrhizal (EM) fungi are ubiquitous in temperate and boreal forests, comprising over 20,000 species forming root symbiotic associations with Pinaceae and woody angiosperms. As much as 100 different EM fungal species can coexist and interact with the same tree species, forming complex multispecies networks in soils. The degree of host specificity and structural properties of these interaction networks (e.g., nestedness and modularity) may influence plant and fungal community assembly and species coexistence, yet their structure has been little studied in northern coniferous forests, where trees depend on EM fungi for nutrient acquisition. We used high‐throughput sequencing to characterize the composition and diversity of bulk soil and root‐associated fungal communities in four co‐occurring Pinaceae in a relic foredune plain located at Îles de la Madeleine, Québec, Canada. We found high EM fungal richness across the four hosts, with a total of 200 EM operational taxonomic units (OTUs), mainly belonging to the Agaricomycetes. Network analysis revealed an antinested pattern in both bulk soil and roots EM fungal communities. However, there was no detectable modularity (i.e., subgroups of interacting species) in the interaction networks, indicating a low level of specificity in these EM associations. In addition, there were no differences in EM fungal OTU richness or community structure among the four tree species. Limited shared resources and competitive exclusion typically restrict the number of taxa coexisting within the same niche. As such, our finding of high EM fungal richness and low host specificity highlights the need for further studies to determine the mechanisms enabling such a large number of EM fungal species to coexist locally on the same hosts.     

Strong host preference of ectomycorrhizal fungi in a Tasmanian wet sclerophyll forest as revealed by DNA barcoding and taxon-specific primers

Ectomycorrhizal (ECM) symbiosis is a widespread plant nutrition strategy in Australia, especially in semiarid regions. This study aims to determine the diversity, community structure and host preference of ECM fungi in a Tasmanian wet sclerophyll forest. Ectomycorrhizal fungi were identified based on anatomotyping and rDNA internal transcribed spacer (ITS)-large subunit (LSU) sequence analysis using taxon-specific primers. Host tree roots were identified based on root morphology and length differences of the chloroplast trnL region. A total of 123 species of ECM fungi were recovered from root tips of Eucalyptus regnans (Myrtaceae), Pomaderris apetala (Rhamnaceae) and Nothofagus cunninghamii (Nothofagaceae). The frequency of two thirds of the most common ECM fungi from several lineages was significantly influenced by host species. The lineages of Cortinarius, Tomentella-Thelephora, Russula-Lactarius, Clavulina, Descolea and Laccaria prevailed in the total community and their species richness and relative abundance did not differ by host species. This study demonstrates that strongly host-preferring, though not directly specific, ECM fungi may dominate the below-ground community. Apart from the richness of Descolea, Tulasnella and Helotiales and the lack of Suillus-Rhizopogon and Amphinema-Tylospora, the ECM fungal diversity and phylogenetic community structure is similar to that in the Holarctic realm.     

Demographic and environmental drivers of metagenomic viral diversity in vampire bats

Viruses infect all forms of life and play critical roles as agents of disease, drivers of biochemical cycles and sources of genetic diversity for their hosts. Our understanding of viral diversity derives primarily from comparisons among host species, precluding insight into how intraspecific variation in host ecology affects viral communities or how predictable viral communities are across populations. Here we test spatial, demographic and environmental hypotheses explaining viral richness and community composition across populations of common vampire bats, which occur in diverse habitats of North, Central and South America. We demonstrate marked variation in viral communities that was not consistently predicted by a null model of declining community similarity with increasing spatial or genetic distances separating populations. We also find no evidence that larger bat colonies host greater viral diversity. Instead, viral diversity follows an elevational gradient, is enriched by juvenile‐biased age structure, and declines with local anthropogenic food resources as measured by livestock density. Our results establish the value of linking the modern influx of metagenomic sequence data with comparative ecology, reveal that snapshot views of viral diversity are unlikely to be representative at the species level, and affirm existing ecological theories that link host ecology not only to single pathogen dynamics but also to viral communities.     

东北红松纯林菌根外生菌根真菌群落与环境因子的相关性

外生菌根是木本植物根系与真菌形成的共生结构,外生菌根真菌在红松等外生菌根树种的定植与森林生态系统的保持方面起到至关重要的作用。明确菌根系统内外生菌根真菌群落组成是揭示菌根共生机制的前提条件。本研究利用Illumina Hiseq测序平台对生长季内红松纯林内根围土壤及菌根样品ITS2区进行高通量测序,分析其外生菌根真菌群落结构随季节的变化规律,同时通过统计学的方法分析了红松根系微生态中外生菌根真菌群落结构组成变化与其他生物因素、非生物因素的相关性。结果如下：（1）从6月份到10月份,5个月的菌根样品测序共得到741个真菌OTUs,利用FUNGuild数据库分析,其中85个OTUs归类为外生菌根真菌,优势属（相对丰度>5）为蜡壳菌属Sebacina、乳牛肝菌属Suillus、Meliniomyces、红菇属Russula、棉革菌属Tomentella、须腹菌属Rhizopogon和缘腺革菌属Amphinema。6月份菌根中外生菌根真菌的多样性最大,显著高于其他月份。（2）红松林外生菌根真菌群落组成受到土壤pH、有效磷含量、有效钾含量和土壤有效氮含量的影响,它们与外生菌根真菌优势属相对丰度呈现正相关或负相关。（3）根围土壤内真菌是影响红松根系外生菌根真菌相对丰度的另一重要因素,其中,包括普可尼亚属Pochonia、产丝齿菌属Hyphodontia、镰刀菌属Fusarium、Collembolispora、枝穗霉属Clonostachys、Apodus、鹅膏属Amanita在内的土壤真菌与根内外生菌根真菌的相对丰度呈线性关系。同时,超过85%的根内外生菌根真菌与同一取样地的土壤共有,可以认为侵染和扩散是红松根内外生菌根真菌群落形成的主要方式,同时兼有植物根系的选择,因为根内并不包括所有土壤中存在的外生菌根真菌,其机制需要进一步人工模拟试验验证。     

Changes in fungal community of Scots pine (Pinus sylvestris) needles along a latitudinal gradient in Sweden

Conifer needles are typically long lived and can host a diverse community of fungal species with various effects on their host tree. The purpose of this study was to analyse shifts in the fungal community of Scots pine (Pinus sylvestris) needles on different spatial scales using 454 pyrosequencing. The fungal community composition changed gradually along a north-south gradient through Sweden, representing boreal to temperate vegetation zones. OTU richness and Shannon's diversity index increased with increasing latitude, but only in naturally regenerated forests. On the tree level, needles with symptoms of disease hosted a more diverse mycobiota compared to healthy needles, presumably supporting more pathogenic or saprotrophic species. This study provides a better insight into the patterns of fungal communities of Scots pine needles and highlights the need for further experimental research to identify specific environmental factors shaping the abundance of different fungal species.     

Biogeographic and phylogenetic patterns in diversity of liverwort-associated endophytes

Liverworts harbor diverse fungi, including endophytes, in their healthy tissues. To address whether patterns of endophyte diversity are correlated with host phylogeny or geography, we designed a broad geographic survey with controlled phylogenetic host sampling. We collected liverworts in North Carolina, Washington, Idaho, British Columbia, Germany, and New Zealand and identified endophytes using culture-based and molecular methods. Of the major lineages of filamentous ascomycetes recovered, 53-88% belonged to the Xylariales. Endophyte accumulation curves did not saturate, and singleton sequences were dominant in each region, suggesting that liverwort endophyte communities are diverse. There was no significant difference in species richness between regional endophyte communities; however, total richness estimators indicated that North Carolina and New Zealand have richer communities than do Germany and the Pacific Northwest. This pattern reflects lower per-host endophyte density and prevalence of a common, shared sequence group in Germany and the Pacific Northwest. Although species richness was relatively low in the Pacific Northwest, the greatest phylogenetic diversity of endophytes was recovered there. Tests for regional and host specificity revealed that endophyte floras of hosts within a geographic area are more similar to one another than to those of closely related hosts. Geographic distance, not host phylogeny, best explains differences among communities.     

Arbuscular mycorrhizal communities in tropical forests are affected by host tree species and environment

Arbuscular mycorrhizal (AM) fungi are mutualists with plant roots that are proposed to enhance plant community diversity. Models indicate that AM fungal communities could maintain plant diversity in forests if functionally different communities are spatially separated. In this study we assess the spatial and temporal distribution of the AM fungal community in a wet tropical rainforest in Costa Rica. We test whether distinct fungal communities correlate with variation in tree life history characteristics, with host tree species, and the relative importance of soil type, seasonality and rainfall. Host tree species differ in their associated AM fungal communities, but differences in the AM community between hosts could not be generalized over life history groupings of hosts. Changes in the relative abundance of a few common AM fungal species were the cause of differences in AM fungal communities for different host tree species instead of differences in the presence and absence of AM fungal species. Thus, AM fungal communities are spatially distinguishable in the forest, even though all species are widespread. Soil fertility ranging between 5 and 9 Mg/ha phosphorus did not affect composition of AM fungal communities, although sporulation was more abundant in lower fertility soils. Sampling soils over seasons revealed that some AM fungal species sporulate profusely in the dry season compared to the rainy season. On one host tree species sampled at two sites with vastly different rainfall, relative abundance of spores from Acaulospora was lower and that of Glomus was relatively higher at the site with lower and more seasonal rainfall.     

Spatial patterns of fungal endophytes in a subtropical montane rainforest of northern Taiwan

Fungal endophytes of plants are ubiquitous and important to host plant health. Wood-inhabiting and foliar endophyte communities from multiple tree hosts were sampled at multiple spatial scales across the Fushan forest dynamics plot in northern Taiwan, using culture-free, community DNA amplicon sequencing methods. Fungal endophyte communities were distinct between leaves and wood, but the mycobiomes were highly variable across and within tree species. Despite this, host tree species was an important predictor of mycobiome community-composition. Within a single common tree species, “core” mycobiomes were characterized using co-occurrence analysis. The spatial co-occurrence patterns of these few species of fungal endophytes appear to explain the strong host effect. For wood endophytes, a consistent core mycobiome coexisted with the host across the extent of the study. For leaf endophytes, the core fungi resembled a more dynamic, “gradient” model of the core microbiome, changing across the topography and distance of the study.     

Strong coupling of plant and fungal community structure across western Amazonian rainforests

The Amazon basin harbors a diverse ecological community that has a critical role in the maintenance of the biosphere. Although plant and animal communities have received much attention, basic information is lacking for fungal or prokaryotic communities. This is despite the fact that recent ecological studies have suggested a prominent role for interactions with soil fungi in structuring the diversity and abundance of tropical rainforest trees. In this study, we characterize soil fungal communities across three major tropical forest types in the western Amazon basin (terra firme, seasonally flooded and white sand) using 454 pyrosequencing. Using these data, we examine the relationship between fungal diversity and tree species richness, and between fungal community composition and tree species composition, soil environment and spatial proximity. We find that the fungal community in these ecosystems is diverse, with high degrees of spatial variability related to forest type. We also find strong correlations between α- and β-diversity of soil fungi and trees. Both fungal and plant community β-diversity were also correlated with differences in environmental conditions. The correlation between plant and fungal richness was stronger in fungal lineages known for biotrophic strategies (for example, pathogens, mycorrhizas) compared with a lineage known primarily for saprotrophy (yeasts), suggesting that this coupling is, at least in part, due to direct plant–fungal interactions. These data provide a much-needed look at an understudied dimension of the biota in an important ecosystem and supports the hypothesis that fungal communities are involved in the regulation of tropical tree diversity.     

Deterministic processes have limited impacts on foliar fungal endophyte communities along a savanna-forest successional gradient

Patterns and drivers of succession provide insight into the mechanisms that govern community assembly, but remain poorly understood for microbial communities. We assess whether successional trends of trees are mirrored by foliar endophyte communities of three tree species across a deterministic woody successional gradient. Additionally, we test the relative contribution of abiotic predictors, biotic factors, and spatial distance between sites in predicting composition and richness of endophyte communities. Unlike the tree community, endophyte communities showed no consistent evidence of deterministic succession. Host identity was the most important factor structuring endophyte community composition; within hosts, spatial distance from the indigenous forest and between samples was important, while environmental predictors had small and inconsistent effects. Much variation in endophyte composition remained unexplained. In contrast, endophyte richness was well-explained by predictor variables. Host identity was most important in predicting endophyte richness, while the effect of other predictors on richness differed between host species. We conclude that deterministic succession in trees did not result in deterministic succession in endophyte communities; instead community assembly was most strongly influenced by host identity; while within hosts, neutral processes may be more important for endophyte assembly than deterministic factors.     

Predicting the similarity of parasite communities in freshwater fishes using the phylogeny,ecology and proximity of hosts

Parasite communities tend to be dissimilar in hosts that are geographically, phylogenetically, ecologically and developmentally distant from one another. The decay of community similarity is a powerful and increasingly common method of studying parasite beta diversity, but most studies have examined only a single type of distance. Here, we evaluate distances based on the phylogeny, ecology, spatial proximity and size of hosts, as predictors of the similarity of parasite communities in individual hosts, host populations and host species. We surveyed parasites in six species of fish collected simultaneously from six localities in the St. Lawrence River, Canada, and species in a common group of larval parasites were discriminated using DNA sequences from barcode region of cytochrome c oxidase I. Distances based on the habitat use patterns of host species were good predictors of short‐term, ecological similarity of parasite communities, such as that operating at the scale of the individual host. The genetic distance between host species was associated with almost all types of similarity at all scales, particularly qualitative and phylogenetic similarity of parasite communities at the level of populations and meta‐populations of hosts. The trophic level, diet, spatial proximity and size of hosts were poor predictors of parasite community similarity. The increased taxonomic resolution provided by molecular data increased the explanatory power of regression models, and different factors were implicated when parasite species were distinguished with DNA barcodes than when larval parasites were lumped into morphospecies, as is commonly practiced.     

Ectomycorrhizal fungal diversity and saprotrophic fungal diversity are linked to different tree community attributes in a field‐based tree experiment

Exploring the link between above‐ and belowground biodiversity has been a major theme of recent ecological research, due in large part to the increasingly well‐recognized role that soil microorganisms play in driving plant community processes. In this study, we utilized a field‐based tree experiment in Minnesota, USA, to assess the effect of changes in plant species richness and phylogenetic diversity on the richness and composition of both ectomycorrhizal and saprotrophic fungal communities. We found that ectomycorrhizal fungal species richness was significantly positively influenced by increasing plant phylogenetic diversity, while saprotrophic fungal species richness was significantly affected by plant leaf nitrogen content, specific root length and standing biomass. The increasing ectomycorrhizal fungal richness associated with increasing plant phylogenetic diversity was driven by the combined presence of ectomycorrhizal fungal specialists in plots with both gymnosperm and angiosperm hosts. Although the species composition of both the ectomycorrhizal and saprotrophic fungal communities changed significantly in response to changes in plant species composition, the effect was much greater for ectomycorrhizal fungi. In addition, ectomycorrhizal but not saprotrophic fungal species composition was significantly influenced by both plant phylum (angiosperm, gymnosperm, both) and origin (Europe, America, both). The phylum effect was caused by differences in ectomycorrhizal fungal community composition, while the origin effect was attributable to differences in community heterogeneity. Taken together, this study emphasizes that plant‐associated effects on soil fungal communities are largely guild‐specific and provides a mechanistic basis for the positive link between plant phylogenetic diversity and ectomycorrhizal fungal richness.     

Diversity and community structure of ectomycorrhizal fungi in a wooded meadow

Wooded meadows are seminatural plant communities that support high diversity of various taxa. Due to changes in land use, wooded meadows have severely declined during the last century. The dominant trees in wooded meadows acquire mineral nutrients via ectomycorrhizal fungi. Using anatomotyping and sequencing of root tips, interpolation and extrapolation methods, we studied the diversity and community structure of ectomycorrhizal fungi in two soil horizons of both managed and forested parts of a wooded meadow in Estonia. Species of Thelephoraceae, Sebacinaceae and the genus Inocybe dominated the whole ectomycorrhizal fungal community of 172 observed species. Forested and managed parts of the wooded meadow harboured different communities of ectomycorrhizal fungi, whereas soil horizon had a negligible effect on the fungal community composition. Diverse soil conditions and host trees likely support the high richness of ectomycorrhizal fungi in the wooded meadow ecosystem. Direct sequencing integrated with interpolation and extrapolation methods are promising to identify the fungi at the species level and to compare species richness between communities of ectomycorrhizal fungi.     

Host effects on ectomycorrhizal fungal communities: insight from eight host species in mixed conifer-broadleaf forests

To advance our understanding of host effects on the community structure of ectomycorrhizal fungi (EMF), EMF communities were compared among different host species, genera and families in two mixed conifer-broadleaf forests in Japan. Using molecular identification methods we examined EMF root tips of eight coexisting species belonging to six genera (three families): Abies and Tsuga (Pinaceae), Betula and Carpinus (Betulaceae) and Fagus and Quercus (Fagaceae). In total, 205 EMF species were detected, and the total richness was estimated to exceed 300 species using major estimators. Of the 55 EMF species occurring three or more times, eight showed significantly biased host preference. A Mantel test showed a significantly negative correlation between EMF community similarity and host taxonomic distance. Detrended correspondence analysis separated EMF communities mainly by host taxonomic and successional status. Thus, EMF communities are similar on hosts with similar taxonomic and successional status. A significant proportion of EMF exhibited host specificity, which may contribute to the extremely diverse EMF community in conifer-broadleaf forests.     

Geographical distribution and diversity of bacteria associated with natural populations of Drosophila melanogaster

Drosophila melanogaster is one of the most widely used model systems in biology. However, little is known about its associated bacterial community. As a first step towards understanding these communities, we compared bacterial 16S rRNA gene sequence libraries recovered from 11 natural populations of adult D. melanogaster. Bacteria from these sequence libraries were grouped into 74 distinct taxa, spanning the phyla Proteobacteria, Bacteroidetes, and Firmicutes, which were unevenly spread across host populations. Summed across populations, the distribution of abundance of genera was closely fit by a power law. We observed differences among host population locations both in bacterial community richness and in composition. Despite this significant spatial variation, no relationship was observed between species richness and a variety of abiotic factors, such as temperature and latitude. Overall, bacterial communities associated with adult D. melanogaster hosts are diverse and differ across host populations.     

Contrasting ectomycorrhizal fungal communities on the roots of co-occurring oaks (Quercus spp.) in a California woodland

Plant host species is considered an important factor influencing ectomycorrhizal (EM) communities. To gain insights into the role of host species in structuring EM communities, EM communities on sympatric oak (Quercus) species were compared in the Sierra Nevada foothills of California. Using molecular methods (polymerase chain reaction, cloning, restriction fragment length polymorphism and DNA sequencing), EM fungi on roots of deciduous Quercus douglasii and evergreen Quercus wislizeni trees were identified from 64 soil cores. The total EM species richness was 140, of which 40 taxa were detected on both oak hosts. Greater diversity and frequency of EM fungi with epigeous fruiting habit were found on Q. wislizeni, while taxa in the Ascomycota were more frequent and diverse on Q. douglasii. Using ordination, it was determined that both soil extractable phosphorus and oak host species explained a significant proportion of the variation in EM species distribution. These results indicate that plant host species can be an important factor influencing EM fungal community composition, even within congeneric trees.     

Fungal diversity in ectomycorrhizal communities: sampling effort and species detection

A number of recent review articles on ectomycorrhizal (ECM) fungal community diversity have highlighted the unprecedented increase in the number of publications on this ecologically important but neglected area. The general features of these species-rich, highly dynamic and complex communities have been comprehensively covered but one aspect crucial to our assessment of diversity, namely the sampling of ECM communities has received less attention. This is a complex issue with two principal components, the physical sampling strategy employed and the life cycle traits of the ECM fungi being examined. Combined, these two components provide the image that we perceive as ECM diversity. This contribution will focus primarily on the former of these components using a recent study from a pine forest in central Sweden to highlight some sampling problems and also to discuss some features common to ECM communities. The two commonly used elements of diversity, species richness and community evenness, present rather different problems in the assessment of ECM diversity. The applicability of using current measures of abundance (number or percentage of root tips colonised) to determine community evenness is discussed in relation to our lack of knowledge on the size of individual genets of ECM fungi. The inherent structure of most ECM communities, with a few common species and a large number of rare species, severely limits our ability to accurately assess species richness. A discussion of theoretical detection limits is included that demonstrates the importance of the sampling effort (no. of samples or tips) involved in assessing species richness. Species area abundance plots are also discussed in this context. It is suggested that sampling strategy (bulk samples versus multiple collections of single tips) may have important consequences when sampling from communities where root tip densities differ. Finally, the need for studies of the spatial distribution of ECM on roots in relation to small-scale soil heterogeneity and of the temporal aspects of ECM community dynamics is raised.     

Diversity,distribution and host-species associations of epiphytic orchids in Nepal

Associations between epiphytes and their hosts are among the main factors affecting the biodiversity and distribution of epiphytes. While several previous studies explored the association between epiphyte diversity and host characteristics, very little is known about the generality of such associations at larger spatial scales. We aim to explore the associations between diversity and distribution of epiphytic orchids and host characteristics in different localities in Nepal. Epiphytic orchids and their hosts were recorded along the transects in total of 23,539 host individuals. To describe the diversity of orchids in the different localities, a rarefaction function was used. Univariate and multivariate analyses were carried out to explore the associations with locality, host characteristics, and their interactions with locality. In total, we recorded 141 species of orchids growing on 192 host species. The five localities significantly differed in orchid diversity and abundance. The number of orchid individuals per host species significantly increased with increasing number of host individuals. Species richness, abundances, occupancy of orchid species on host species and composition of orchids varied across the localities. Species richness and abundance was significantly higher on hosts in the higher strata and differed between families of host species. Abundance was higher on evergreen hosts. Composition of orchid communities are also associated with host characteristics, such as habit (shrub/tree/climber), bark texture, nature (deciduous/evergreen) and the plant families of host species. This study revealed a high diversity of epiphytic orchids in the localities studied and strong associations between the orchids and their hosts. Future studies looking at the relationships between epiphyte communities and host characteristics need to identify relationships at a wider scale in order to determine whether they are really general rather than site-specific.